Acta zoologica cracoviensia, 47(3-4): , Kraków, 31 December, 2004

Acta zoologica cracoviensia, 47(3-4): 231-248, Kraków, 31 December, 2004 Mayflies of the Crimean Peninsula. III. The description of Baetis milani sp. n. with notes on taxonomy of the subgenus Rhodobaetis JACOB, 2003 (Ephemeroptera: Baetidae) Roman J. GODUNKO, Grigorii A. PROKOPOV and Tomáš SOLDÁN Received: 18 June, 2004 Accepted for publication: 22 Sept., 2004 GODUNKO R. J., PROKOPOV G. A., SOLDÁN T. 2004. Mayflies of the Crimean Peninsula. III. The description of Baetis milani sp. n. with notes on taxonomy of the subgenus Rhodobaetis JACOB, 2003 (Ephemeroptera: Baetidae). Acta zoologica cracoviensia, 47(3-4): 231-248. Abstract. Baetis (Rhodobaetis) milani sp. n. (larva, as well as reared male and female subimago and imago) is described from numerous localities in foothills and mountains of the Peninsula. Critical distinguishing characters are illustrated and available data on biology and distribution of the new species are summarized. Taxonomic characters of West Palaearctic species of the subgenus Rhodobaetis JACOB, 2003 are discussed in detail. Key words: Ephemeroptera, Baetidae, Baetis, Rhodobaetis, new species, Crimean Peninsula. Roman J. GODUNKO, State Museum of Natural History, National Academy of Sciences of Ukraine, Teatral na 18, UA-79008 L viv, Ukraine. E-mail: godunko@museum.lviv.net; godunko@seznam.cz Grigorii A. PROKOPOV, Tavrida National University V. I. Vernads kogo, Faculty of Geography, Department of Geoecology, Vernadsky Prosp. 4, UA-95007 Simferopol, Ukraine. E-mail: prokopov@crimea.com Tomáš SOLDÁN, Institute of Entomology, Academy of Sciences of the Czech Republic and Biological Faculty, University of South Bohemia, Branišovská 31, CZ-370 05 Èeské Budìjovice, Czech Republic. E-mail: soldan@entu.cas.cz I. INTRODUCTION This paper is the third part of a series of works which deal with mayflies of the Crimean Peninsula, namely with species of the genus Baetis LEACH, 1815 (the subgenus Rhodobaetis JACOB, 2003). The first species of Rhodobaetis from the Crimea, B. braaschi ZIMMERMANN, 1980 was described from the Southern Coast of the Peninsula (ZIMMERMANN, 1980a) and attributed by the author to rhodani species-group s. MÜLLER-LIEBENAU (1969). Later on, KISELEVA & VASYUTA (1984) and TEMIROVA, PARTOLAKHA & TUROBOV (1984) recorded B. rhodani in rivers discharging into Simferopol Reservoir and in to the Biyuk-Karasu River. In following works this taxon was cited from northern slopes of the Crimean Mountains (see KISELEVA, 1993, 1997). Information on availability of B. stipposus KLUGE, 1982 in the Crimea, which had been previously described from Central Asia by KLUGE (1982), was published by NOVIKOVA (1987). This species was repeatedly cited by KISELEVA (1987, 1992, 1997) from foothill sections of the Crimean rivers. GODUNKO &

232 R. J. GODUNKO et al. PROKOPOV (2003) have described B. rhodani tauricus GODUNKO &PROKOPOV, 2003 both on the basis of their own material and G. A. KISELEVA s collection. The nominal subspecies B. rhodani rhodani PICTET, 1843 has not been found in the Crimea during many-year investigations. Finally, GODUNKO et al. (2004) have published the redescription of little known species B. braaschi with the depiction of larva and imago male in detail, and confirmed its synonymy with B. stipposus. The present paper deals with a description of adults and larvae of a new species found in the Crimean Peninsula. The bionomics and distribution of this species are briefly stated, range of subgenus Rhodobaetis is discussed. Main morphological characters of male imagines and larvae of West Palaearctic species of Rhodobaetis have been presented and compared. A c k n o w l e d g m e n t s.we are sincerely indebted to Milan PUTZ for the material kindly placed at our disposal and to Iryna B. KONOVALOVA for help in English version of the paper. This study has been carried out owing to the cooperation between the Academy of Sciences of Czech Republic and the National Academy of Sciences of Ukraine. II. SYSTEMATICS Baetis milani sp. n. (Figs 1-28) Baethis [sic!] rhodani RICTET [sic!]: KISELEVA &VASYUTA, 1984: 143 (partim) Baetis rhodani: TEMIROVA, PARTOLAKHA &TUROBOV, 1984: 137; KISELEVA, 1993: 163 (partim) Baethis [sic!] rhodani MULL [sic!]: KISELEVA &EZERNITSKII, 1985: 112 (partim) Baethis [sic!] rhodani: KISELEVA &VASYUTA, 1986: 57 (partim) Baethis [sic!] gr. rhodani: KISELEVA, 1997: 39 (partim) M a t e r i a l e x a m i n e d. Holotype: male imago, Ukraine, Autonomous Republic of the Crimea, small right tributary of Guva River, Uch-Kosh gap, 21.VI.2002, leg. G. A. PROKO- POV. Paratypes: 7 male imagines, 11 male subimagines, 4 female imagines, 1 female subimago, from the same locality and date as holotype, leg. G. A. PROKOPOV; 2 female imagines, Ukraine, Autonomous Republic of the Crimea, middle part of Uchan-Su River, 13.IV.2002, leg. G. A. PROKOPOV; 2 male imagines, 2 male subimagines, 6 female imagines, Ukraine, Autonomous Republic of the Crimea, Kyzyl-Kobinka River (150 m downstream of Hidropost ), 14.VI.2002, leg. G. A. PROKOPOV. Other specimens (not type). Ukraine, Autonomous Republic of the Crimea: 8 larvae, upper part of Al ma River, 12.IV.1984, leg. G. A. KISELEVA; 9 larvae, 1 larval skin, Eastern Ulu-Uzen River, 16.VII.1989, leg. KUZNETSOVA & LYSINKINA; 13 larvae, Chorna River, 16.VII.1989, leg. KUZNET- SOVA & LYSINKINA; 32 larvae, upper part of Guva River, 16.VI.1998, leg. G. A. PROKOPOV; 20 larvae, down part of Voron River, 16.VII.1998, leg. G. A. PROKOPOV; 3 larvae, middle part of Shelen River, 20.VII.1998, leg. G. A. PROKOPOV; 26 larvae, Khasta-Bash River, 31.V.1999, leg. R. J. GODUNKO; 64 larvae, upper part of Kyzyl Koba River near Pereval ne village, 2.VI.1999, leg. R. J. GODUNKO; 37 larvae, upper part of Burul cha River, W slope of Yuki-Tepe Mt., 3.VI.1999, leg. R. J. GODUNKO; 9 larvae, Uchan-Su River (near Uchan-Su waterfall), 17.IX.1999, leg. G. A. PROKO- POV; 53 larvae, Guva River (Uch-Kosh gorge), 20.IX.1999, leg. G. A. PROKOPOV; 14 larvae, upper part of Guva River, 24.V.2000, leg. G. A. PROKOPOV; 21 larvae, Voron River, 4.VI.2000, leg. G. A. PROKOPOV; 34 larvae, down part of Shelen River, 7.VI.2000, leg. G. A. PROKOPOV; 65 larvae, down pat of Eastern Ulu-Uzen River, 11.VI.2000, leg. G. A. PROKOPOV; 73 larvae, stream near Pryvitne village, 18.VI.2000, leg. R. J. GODUNKO & M. PUTZ; 24 larvae, Khasta-Bash River, 20.VI.2000, leg. R. J. GODUNKO &M.PUTZ; 2 male subimagoes, 2 female imagoes, 2 females subimagoes, 64 larvae, middle part of Eastern Ulu-Uzen River, 28.VII.2000, leg. G. A. PROKOPOV; 1 female imago, 1 female subimago, 25 larvae, Eastern Ulu-Uzen River near Solnechnogorskoe village, 28.VII.2000, leg. G. A. PROKOPOV; 13 larvae, upper part of Eastern Ulu-Uzen River (near water-

Mayflies of the Crimean Peninsula 233 falls), 15.X.2000, leg. G. A. PROKOPOV; 9 larvae, Eastern Ulu-Uzen River near General s ke village, 15.X.2000, leg. G. A. PROKOPOV; 28 larvae, Eastern Ulu-Uzen River near General s ke village, 2.V.2001, leg. G. A. PROKOPOV; 46 larvae, 2 female imago, middle part of Eastern Ulu-Uzen River, 2.V.2001, leg. G. A. PROKOPOV; 1 male imago, 12 larvae, Al ma River (higher than a trout facilities), 28.VI.2001, leg. G. A. PROKOPOV; 2 larvae, Al ma River (near trout facilities), 28.VI.2001, leg. G. A. PROKOPOV; 38 larvae, 1 larval skin, Al ma River near Asport Boundary, 29.VI.2001, leg. G. A. PROKOPOV; 18 larvae, Al ma River near Sosnovyi Boundary, 1.VII.2001, leg. G. A. PROKOPOV; 11 larvae, Chorna River (source of the river near Rodnikovskoe village), 9.VII.2001, leg. G. A. PROKOPOV; 2 larvae, Alaka River near Dzhurla waterfall, 18.VII.2001, leg. G. A. PROKO- POV; 54 larvae, Uzundzha River (upper then Kolhoznoe village), 2.V.2002, leg. G. A. PROKOPOV; 7 larvae, Uzundzha River (upper then Rodnikovskoe village), 3.V.2002, leg. G. A. PROKOPOV. All adults of type series were reared from larvae. The specimens were preserved in 80% alcohol. All larval skins of type specimens are mounted on microscopic slide. The holotype and some paratypes are housed in the collection of the State Museum of Natural History, National Academy of Sciences of Ukraine (L viv). Some other paratypes can be found in the Institute of Entomology, Academy of Sciences of the Czech Republic and in the collection of G. A. PROKOPOV. Description. Male imago. Length: body 7.0-7.6 mm; fore wings 5.9-6.7; cerci 13.0-14.0 mm; length of fore legs: femora + tibiae = 2.9-3.0 mm; tarsal segments: T1 = 0.67-0.75 mm; T2 = 0.65-0.67 mm; T3 = 0.35-0.37 mm; T4 = 0.20-0.25 mm; general ratio of tarsal segments length: 1>=2>3>4 (Fig. 3). General color of body light, yellow to light brown. Head yellow to light brown. Antennae light brown. Flagellum slightly darker proximally. Ocelli black at the base and yellowish. Eyes surrounded by a yellowish ring. Turbinate eyes oval in dorsal view with convex external margin (Fig. 2). Facetted surface of turbinate eyes orange, sometimes surrounded by a brownish ring. The shaft lighter than facetted surface, yellow or yellowish-brown. Basal part of turbinate eyes shaft with apparently narrow orange or light brown ring (Fig. 1). Thorax light, yellow to light-brown, with some brownish maculation dorsally and laterally. Pronotum with dark smudges centrally. Dorsal part of thorax with longitudinal light brown strip. Metanotum darker than pro- and mesonotum, brownish color. Fore wings hyaline, transparent. Pterostigma only slightly opaque, with 5-9 simple cross veins, some of them with pronounced bifurcation. Venation yellowish to light brown. Hind wings hyaline, transparent, with three simple longitudinal veins and distinct costal process typical of subgenus Rhodobaetis (Fig. 4). Legs light, yellow to yellowish-brown. Femora and tibia of fore legs light, yellow. Femora with hardly visible smudges distally. Tibiae slightly darker distally. Tarsal segments yellowish-brown, segment 4 darker. Middle and hind legs the lightest, uniformly yellow. Abdominal tergum I darker than others. Terga II-X uniformly yellowish-brown to light brown. Occasionally terga VII-X slightly darker. Posterior margin of terga sometimes with narrow reddish-brown band. Lateral part of terga with brownish smudges. Sterna the same color that terga, with small unclear spots. Cerci yellowish-white to yellowish-brown, slightly darker at the base. Joints of caudal segments dark. Genitalia yellow to light brown. Basal segment of forceps nearly as long as wide (Figs 5, 6). Segment 1 relatively wide at the base with subparallel margins. Segment 2 elongated and narrow, widening towards segment 3, the widened distal part being as long as 1/2 the segment length. Inner margin of segment 2 slightly concave. Segment 3 of various structure, oval or slightly elongated, with more or less truncate inner margin (Figs 5, 6). Female imago. Length: body 8.0-8.5 mm; fore wings 7.8-8.3; cerci 10.0-11.0 mm. General color of body yellow to yellowish-brown, sometimes distinctly brown, darker than in male imago. Head uniformly yellow or light brown. Antennae with brownish flagellum, darker than head. Eyes and base of ocelli black; apical part of ocelli whitish. Thorax with brownish bands dorsally and laterally. Wings hyaline, transparent. Pterostigma slightly milky color. Venation light brown to brown. Legs uniformly yellow to yellowish-brown. Fore legs slightly darker than middle and hind ones. All femora distally with diffuse smudges. Distal part of tibiae and tarsi darker than femora. Abdominal terga yellow to yellowish-brown, with

234 R. J. GODUNKO et al. Figs 1-4. Baetis milani sp. n., male imago: 1 head (lateral view); 2 head (dorsal view); 3 fore leg; 4 hind wing. brownish band near posterior margin. Sterna lighter than terga, with small spots centrally. Cerci yellow to yellowish-brown. Male subimago. Length: body 6.4-7.2 mm; fore wings 5.6-6.0; cerci 9.1-11.2 mm. General color of body yellowish-brown to light brown with light greyish ting. Head yellow to yellowish-brown. Antennae generally the same color that head, flagellum slightly darker. Eyes and basal part of ocelli black. Ocelli whitish. Facetted surface of turbinate eyes light orange, sometimes with narrow marginal ring. The shaft lighter, yellow, with well visible narrow brownish ring basally. Thorax yellowish-brown to light brown with brown or black bands. Wings uniformly yellowish-grey. Venation yellowish. Legs yellowish-brown. Fore legs the same color that middle and hind legs. All femora with clear dark spots distally. Joints of leg segments dark.

Mayflies of the Crimean Peninsula 235 Figs 5-6. Baetis milani sp. n., male imago: 5-6 genitalia (ventral view). Abdominal terga yellowish-brown, with some small central spots and L-shaped light spots laterally. Sterna the same color that terga, with light spots near anterior part of segments. Cerci uniformly yellowish-brown. Female subimago. Length: body 6.8-8.4 mm; fore wings 7.4-7.6; cerci 8.2-8.7 mm. General color of body similar to male subimago, somewhat lighter. Mature larva. Length of body: 7.5-8.5 mm (male larva), 7.7-9.4 mm (female larva); length of cerci: 4.6-6.0 mm; length of terminal filament: 2.9-4.3 mm. General color of body yellowish-brown to brown. Head yellowish-brown, antennae slightly darker. Numerous spatulas on the frons surface with margins convergent (Fig. 7). Surface of larval turbinate eyes dark orange, with yellowish ring around the margin. Pedicel with elongated, sharply pointed or bluntly pointed scales arranged in one irregular row or scattered over the segment surface (Figs 8, 15). Scape with small scales and fine hairs. Labrum distinctly wide (the width/length ratio = 1.42), with 1 + 6-10 (mainly 7-9) long submarginal bristles arranged in one regular or irregular row (Fig. 12), occasionally in two rows (Fig. 13). Labrum laterally with few fine bristles. Apical part of segment 2 of maxillary palps with well visible scales on rounded projection and fine hairs (Fig. 9). First tooth of mandibular incisor approximately quadrate, other teeth relatively rounded (Fig. 10). Segment 3 of labial palp rounded, relatively wide, only slightly asymmetric. The length/width ratio of segments 2+3 of labial palps = 2.0 (in B. rhodani rhodani mainly 2.0 (THOMAS & SOLDÁN, 1987); in B. rhodani tauricus is 2.0-2.2 (GODUNKO &PROKOPOV, 2003)) (Fig. 11). Glossae and paraglossae relatively narrow (Figs 18, 19). Paraglossae with three regular rows of bristles at the apex.

236 R. J. GODUNKO et al. Thorax yellowish-brown to brown. Pronotum with a pair of central oval diffuse spots and pair of smudges laterally. Medial longitudinal strip generally present. Mesonotum with yellowish-brown longitudinal bands and light spots near larval wing base. Legs yellowish-brown to brown. Femora with central diffuse large dark spot or with small spot distally. External margin of femora with dense row of fine long setae sharply pointed at the apex and arranged in 2-3 rows proximally (in central Figs 7-13. Baetis milani sp. n., larva: 7 surface of frons; 8 base of antenna; 9 apex of maxillary palps; 10 mandibular incisors; 11 labial palp; 12-13 labrum.

Mayflies of the Crimean Peninsula 237 Figs 14-17. Baetis milani sp. n., larva: 14 posterior margin of femur; 15 surface of pedicel; 16 surface of tergite 4; 17 surface of paraproct plate. Scale bar = 10 m. part of femora in 1-2 rows) (Fig. 21). These setae distinctly thinner and longer than in B. rhodani rhodani and similar to those of B. rhodani tauricus (Fig. 14). Inner margin of femora with small spines. Surface of femora with spines and spatulas, spine bases, hardly visible triangular scales and fine hairs. Tibiae the same color that femora or slightly darker. Tarsi brown, distinctly darker distally. External and inner margin of tibiae and tarsi with spines and fine hairs. Occasionally tarsi with two rows of spines distally (Fig. 20). Tarsal claw brown, relatively elongated, with 9-12 teeth and without hairs. Abdominal terga yellowish-brown to light brown, occasionally dark brown: tergum I generally light, with diffuse central spot; terga II and III generally uniformly color, with two central oval spots and some small spots; tergum IV light, with large central spot and two small spots; tergum V the lightest, only with one central light spot and longitudinal darker spots laterally; color of terga VI-VIII generally similar to terga II and III; terga IX and X light, generally uniformly color, without distinct spots. Medial longitudinal strip well visible on all terga. Lateral part of all terga with broad light spots. Posterior margin of abdominal terga with narrow dark brown strip. Sterna lighter than terga. Posterior margin of abdominal terga of different structure: posterior margin of terga I-V (occasionally terga I-VI) generally with irregular row of sparse spatulas with margins distinctly convergent and with bluntly pointed spatulas alternating with fine hairs only; terga VII and VIII (occasionally tergum IX) with more or less regular row of triangular spines alternating with scales and fine hairs; terga IX and X generally with prevalence of triangular spines (Fig. 22). Surface of terga with sharply pointed or bluntly pointed scales, numerous broad triangular scales and fine hairs (Fig. 16). Gill 1 slightly asymmetric, with hardly visible tracheization (Fig. 24). Its external margin mainly without spines. Gills 2-6 distinctly asymmetric with row of spines on external margin (Fig. 23). Gill 7 apparently symmetrical with spines on external margin (Fig. 25). Paraproct

238 R. J. GODUNKO et al. Figs 18-21. Baetis milani sp. n., larva: 18 glossa; 19 paraglossa; 20 tarsal claw; 21 hind leg. plate with 9-14 teeth on the margin (Figs 27, 28). Surface of paraproct with a few sharply pointed scales, bases of scales and fine hairs (Figs 17, 27). Cerci yellowish-brown to brown, distinctly darker. Terminal filament 3/4 the cerci length. A f f i n i t i e s. B. milani sp. n. clearly belongs to Rhodobaetis and occupies a relatively isolated position among other species of the subgenus. This species can be distinguished by the following combination of characters: in imago male: (I) light color of body; (II) facetted surface of turbinate eyes orange; (III) shaft yellow or yellowish-brown; (IV) segment 2 of forceps elongated and narrow, widening towards segment 3, the widened distal part being as long as 1/2 the segment length; in larva: (V) frons surface with numerous spatulas with margins convergent ; (VI) pedicel with sharply pointed or bluntly pointed scales; (VII) scape with small scales; (VIII) labrum distinctly wide with 1 + 6-10 long bristles arranged in one row (rarely in two rows); (IX) apical part of segment 2 of maxillary palps with well visible scales; (X) segment 3 of labial palp rounded, rela-

Mayflies of the Crimean Peninsula 239 tively wide, only slightly asymmetric; (XI) paraglossae with three regular rows of bristles at the apex; (XII) external margin of femora with dense row of fine long setae sharply pointed at the apex and arranged in 2-3 rows proximally; (XIII) tarsal claw with 9-12 teeth and without hairs ; (XIV) decoration of posterior margins and surface of abdominal terga; (XV) gills 2-6 distinctly asymmetric with row of spines on external margin; (XVI) paraproct plate with 9-14 teeth on the margin and sharply pointed scales on the surface; (XVII) terminal filament 3/4 the cerci length. By Figs 22-28. Baetis milani sp. n., larva: 22 posterior margin of abdominal tergite 4; 23 fourth gill; 24 first gill; 25 seventh gill; 26 external margin of fourth gill; 27 paraproct plate; 28 inner margin of paraproct plate.

240 R. J. GODUNKO et al. Fig. 29 Distribution of B. milani sp. n. in the Crimean Peninsula. combination of larval characters (VI-VIII, XII, XIV-XVI) B. milani sp. n. clearly differs from other species of the subgenus Rhodobaetis. Male imago of a new species markedly differs from other species of the subgenus by the color of turbinate eyes and body. B. milani sp. n. can be easily distinguished from other species of Rhodobaetis from the Crimean Peninsula, viz., B. braaschi and B. rhodani tauricus: from the first one by a number of characters in male imago (II-III) and in larva (VI-X, XII, XIV-XVI); from the latter one by a number of characters in male imago (I-III) and in larva (VI, VII, XIV, XVI). Etymology.Thenewspecies is named in honor of our friend Milan PUTZ, who collected some specimens of this species in the Crimean Peninsula. Distribution and bionomics.b. milani sp. n. inhabits the upper reaches of mountain rivers of the Crimea (Fig. 29). It is especially abundant in the rivers of the Southern Coast, frequently being there the only species of the genus Baetis. In those rivers the species larvae can be found everywhere down the stream nearly to their mouths. Some river sections, where B. milani sp. n. larvae occur, dry up in summer. Water habitats in generally are characterized by fast current, rocky bed with admixture of detritus at 0.2-0.5 m depth at the altitudes between 900 m to 200 m (Figs 30, 31). Water temperature in those localities fluctuates from 6 C to 20 C and mineralization of water between 340 mg/l and 700 mg/l. In chemical composition of these waters, the ions HCO 3 - (177-300 mg/l) and Ca 2+ (44-110 mg/l) predominate. B. milani sp. n. forms a part of a community of epi- and metarhithral of the Crimean rivers, where occurs together with following representatives of aquatic fauna: Electrogena braaschi (SOWA, 1984), Siphonoperla taurica (PICTET, 1841), Agapetus ajpetriensis MARTYNOV, 1917, Plectrocnemia intermedia MARTYNOV, 1917, Silo alupkensis MARTYNOV, 1917, Cnetha spp., Dugesia gonocephala taurocaucasica PORFIRIEVA, 1958 and others. The new species is probably an endemic of the Crimea.

Mayflies of the Crimean Peninsula 241 Figs 30-31. Typical habitat of B. milani sp. n.: 30 Eastern Ulu-Uzen River; 31 Al ma River. III. DISCUSSION The name Rhodobaetis was suggested for the first time by KAZLAUSKAS (1972: 337) for species united by MÜLLER-LIEBENAU (1969: 91) in rhodani species-group. Subgeneric names given by R. S. KAZLAUSKAS, Rhodobaetis in particular, were invalid, since type species were not determined (HUBBARD, 1979). Subsequently, NOVIKOVA & KLUGE (1987) designated type species of some subgenera, previously distinguished by R. S. KAZLAUSKAS, having made their names suitable for application. How-

242 R. J. GODUNKO et al. ever, species which at present belong to Rhodobaetis, were considered by NOVIKOVA & KLUGE (1987: 8) as those belonging to rhodani species-group of the genus Baetis s. str. and this point of view was generally accepted. JACOB (2003: 122) made use of the old name given by R. S. KAZLAUSKAS and made it appropriate having designated the type species of the subgenus Rhodobaetis, viz. B. rhodani rhodani. The diagnosis suggested by JACOB (2003) for the subgenus Rhodobaetis, is based on the characters of larvae and male imagines. The distinguishing character of larvae, according to the author, is the availability of movable spatulas on terga surface. The distinguishing characters of male imagines are as follows (JACOB, 2003): basal segment of forceps with weakly distinct protuberances on inner margin; segment 1 of forceps cylindrical or slightly conical, rather sharply narrowing ; segment 2 of forceps narrow at the base; segment 3 of forceps almost spherical, separated from segment 2 by incision on outer margin; hind wing with three simple longitudinal veins. Larval scales described by JACOB (2003) are really present on terga surface of most species Rhodobaetis, however their shape is quite different, from spatulas widening, as in B. braaschi (see GODUNKO et al. 2004), to pointed scales, as in B. milani sp. n. (Figs 8, 15). And what is more, wide spatulas are also present on terga and paraproct surfaces in the species B. pseudothermicus KLUGE, 1983 (KLUGE, 1983: Figs 11, 12), which has been attributed by NOVIKOVA (1987: 80) to separate pseudothermicus species-group. There are no special differences in structure of genitalia between species of Rhodobaetis and pseudothermicus species-group. The same way as in some species of Rhodobaetis (e. g. in B. rhodani tauricus), B. pseudothermicus is lacking protuberance on inner margin of basal segment of forceps (KLUGE, 1983: Fig. 15, GODUNKO & PROKOPOV, 2003: Fig. 3). Three simple longitudinal veins on hind wings are typical of all discussed groups, just as of Rhodobaetis. SOLDÁN (1977a) has mentioned the availability of only two longitudinal veins in B. baksan SOLDÁN 1977. NOVIKOVA (1987) has noted, that in fact there is the third vein, which is very close to posterior margin of hind wings. Only in one case (undescribed imaginal material of Rhodobaetis from Algeria) a fork second vein is present on hind wing (possible individual variability of specimen). All the facts stated above afford ground to consider the species attributed to pseudothermicus species-group as close to the species of the subgenus Rhodobaetis s. JACOB (2003). Since no significant differences in structure of larvae and male imagines have been revealed, it is advisable to attribute these groups to the subgenus Rhodobaetis. In this connection we suggest to define more precisely and to amplify the earlier diagnoses of rhodani species-group (MÜLLER-LIEBENAU, 1969: 91; MORIHARA & MCCAFFERTY, 1979: 146; NOVIKOVA, 1987: 66-67), and of the subgenus Rhodobaetis (JACOB, 2003: 122). The following combination of cited characters makes possible to distinguish species of Rhodobaetis (incl. B. pseudothermicus) from other species of Baetis s. l.: (I) surface of terga with scales of different structure (a wide, widened spatulas, rounded at the tip; b relatively narrow spatulas with margins slightly divergent, rounded at the tip; c relatively narrow scales with margins slightly convergent, rounded or bluntly pointed at the tip; d scales with wide base and margins distinctly convergent, pointed at the tip); (II) antennal segments (mainly pedicel), surface of femora and paraproct plate with every kind of scales mentioned above; (III) basal segment of forceps approximately square or slightly elongated; (IV) inner margin of basal segment of forceps sometimes with small subapical protuberance; (V) segment 1 of forceps with subparallel margins basally and medially, distinctly or slightly convergent ; (VI) inner margin of segment 1 of forceps with subapical protuberance; (VII) segment 2 of forceps narrow at base, mainly widened towards the apex; (VIII) segment 3 of forceps of various structure (a oval and slightly asymmetric; b almost square with truncate inner margin; c almost spherical; d elongated with length/width ratio = 1.5-3.0). Up to date 22 species of the subgenus Rhodobaetis are known from Palaearctic region (incl. Palaearctic parts of China). The species B. bicaudatus DODDS, 1923 has Holarctic distribution, besides Nearctic occurring from Japan and the Far East to the Arctic Urals. One species, namely B. thermicus UENO, 1931 is known from Japan (FUJITANI, HIROWATARI, TANIDA, 2003). B. pseudothermicus and B. silvaticus KLUGE, 1983 (KLUGE, 1983) have been described from the Primorye Territory. The fauna of Central Asia includes the most number of species of Rhodobaetis. The first Central Asian species of the subgenus, those are B. issyksuvensis BRODSKY, 1930, B. mycetopis BRODSKY,

Mayflies of the Crimean Peninsula 243 1930 and B. heptapotamicus BRODSKY, 1930 were described from Kyrgyzstan and Kazakhstan (BRODSKY, 1930). Later on, some of those species were also found and redescribed by KLUGE (1982) also from Tajikistan and Uzbekistan. B. oreophilus KLUGE, 1982 was described by larvae and associated imago from Kyrgyzstan (KLUGE, 1982). B. rhodani rhodani has Euro-Asian range. The subgenus Rhodobaetis is presented in the Crimea by three species: B. braaschi, B. milani sp. n. and B. rhodani tauricus. B. braaschi, which was regarded earlier the endemic of the Crimean Mountains (ZIMMERMANN, 1980a), belongs to the group of Euro-Asian species and has been recorded also in the Caucasus and all over Central Asia as well (KLUGE, 1982, GODUNKO et al. 2004). Besides the latter, two more, probably endemic species have been found in the Crimea, those are B. rhodani tauricus (GODUNKO &PROKOPOV, 2003) and above described B. milani sp. n. Two species of Rhodobaetis were described from the Southern and Central Caucasus: B. baksan by adults and larvae found in the Central Caucasus and B. ilex JACOB &ZIMMERMANN, 1978 only by larvae from Georgia and Armenia (SOLDÁN, 1977a, JACOB & ZIMMERMANN, 1978). The information of ZIMMERMANN (1980b: 102, fig. 6) on finding of B. gemellus EATON, 1885 in the Caucasus and the Transcaucasus probably regards to a new species of the subgenus Rhodobaetis. The larvae of B. pseudogemellus SOLDÁN, 1977, B. rhodani sinespinosus SOLDÁN & THOMAS, 1883 and B. bisri THOMAS & DIA, 1983 were described from Middle East and Northern Africa (SOLDÁN, 1977b; SOLDÁN &THOMAS, 1983; THOMAS &DIA, 1983). MÜLLER-LIEBENAU (1971) has described two species, those are B. canariensis MÜLLER-LIEBENAU, 1971 and B. pseudorhodani MÜLLER-LIEBENAU, 1971 from the Canary Islands. Larvae of one more species of the subgenus Rhodobaetis, namely B. ingridea THOMAS &SOLDÁN, 1987 were described from the Corsica (THOMAS &SOLDÁN, 1987). THOMAS (1999) having analyzed the material from the Pyrenees, as well as the literature data and the results of many-year research in species of Rhodobaetis in France, Switzerland and Italy has suggested the species name B. gadeai THOMAS, 1999 for B. gemellus s. MÜLLER-LIEBENAU (1969). In THOMAS s opinion that species is the Pyrenees endemic. ENGBLOM (1996) has assumed that designation rhodani embraces the complex of several forms which taxonomic status is unknown. The description of the subspecies B. rhodani tauricus from the Crimea confirms this assumption. The comparative analysis of main morphological characters of larvae and male imagines of West Palaearctic species of the subgenus Rhodobaetis are given in Tables 1-3. Since the validity of the species name of B. gemellus s. EATON, 1885 is called in question, it is absent from the tables given. Tables 1-3 Main morphological characters for distinguishing of West Palaearctic species of the subgenus Rhodobaetis. CH characters; larva: 1 pedicel (shape of scales); 2 scape (shape of scales); 3 labrum (the mean width/length ratio); 4 labrum (number of long submarginal bristles); 5 maxillary palps (apical part of distal segment); 6 paraglossae (number of regular rows of bristles); 7 labial palps; 8 external margin of femora (shape of bristles); 9 tarsal claw (number of strong teeth); 10 tarsal claw (presence of apical setae); 11 surface of terga (shape of scales); 12 posterior margin of terga 3-6 (presence of triangular spines); 13 shape of 3-5 gills; 14 spines of external margin of gills; 15 paraproct plate (number of marginal teeth); 16 paraproct plate (shape of scales); 17 terminal filament length (relative cerci length); male imago: 18 turbinate eyes (facetted surface); 19 shaft of turbinate eyes; 20 coloration of thorax; 21 hind wings (number of veins); 22 basal segment of forceps; 23 segment 1 of forceps; 24 segment 2 of forceps (widened part); 25 segment 2 of forceps (inner margin); 26 segment 3 of forceps. BBA B. baksan; BBI B. bisri; BBR B. braaschi; BC B. canariensis; BG B. gadeai; BIL B. ilex; BIN B. ingridae; BM B. milani sp. n.; BRR B. rhodani rhodani; BRS B. rhodani sinespinosus; BRT B. rhodani tauricus; BPG B. pseudogemellus; BPR B. pseudorhodani.

244 R. J. GODUNKO et al. Table 1 CH BBA BBR BIL BM BRT 1 elongated and pointed elongated and widened elongated and bluntly pointed elongated, pointed or not elongated and bluntly pointed widened 2 small and pointed small and wide mainly absent small and pointed small and wide 3 1.45 1.40 1.16 1.42 1.28 4 1 + 11-14 1 + 5-7 1 + 6-10 1 + 6-10 1 + 6-9 5 with 3-9 short spines with one pointed scale with one rudimentary scale with one pointed scale 6 2-3 3 4-5 3 3 7 slightly asymmetric 8 long and pointed strongly asymmetric and wide short and rounded slightly asymmetric slightly asymmetric and relatively narrow and relatively wide long and pointed long and pointed 9 8-12 8-13 11-18 9-12 7-10 10 absent absent absent absent absent 11 wide and rounded wide and rounded relatively wide and rounded pointed or bluntly pointed with one pointed scale slightly asymmetric and relatively wide long and pointed wide and rounded 12 absent present absent absent present 13 asymmetric strongly asymmetric asymmetric strongly asymmetric slightly asymmetric 14 mainly absent or with 1-3 solitary spines absent present present present 15 2-4 5-10 10-13 9-14 8-14 16 pointed widened pointed pointed widened 17 2/3 2/3-3/4 3/4 3/4 3/4 18 19 ferruginous or dark orange light with ferruginous ring 20 pith-brown light yellow orange orange light yellow, sometimes with dirty yellow ring yellow to yellowish-brown yellow to light, with yellowish-brown with orange-violet ring mainly orange ring yellow to light-yellow 21 3 (2 well visible only) 3 3 3 22 approximately square 23 24 slightly convergent about 1/2 of segment length almost square, slightly convergent with subparallel margins 1/2-2/3 of segment length brown to dark brow approximately square approximately square with subparallel margins 1/2 of segment length 25 slightly concave clearly concave slightly concave 26 oval or slightly elongated oval or asymmetric oval or slightly elongated with subparallel margins 2/3 of segment length straight or slightly concave oval or slightly quadrangular

Mayflies of the Crimean Peninsula 245 Table 2 CH BBI BG BRR BRS BPG 1 elongated and pointed elongated and bluntly pointed elongated, widened or elongated and bluntly pointed pointed elongated, pointed and bluntly pointed 2 elongated and pointed small and wide small and wide mainly absent small and pointed 3 1.24 1.30 1.32 1.38 1.45 4 1 + 5-9 1 + 7-10 1 + 7-12 1 + 9-12 1 + 10-12 5 with one rudimentary scale with one pointed scale with one pointed scale absent with one pointed scale 6 5-6 3 3 4 3 7 relatively slightly asymmetric symmetrical and wide and relatively wide slightly asymmetric and relatively wide slightly asymmetric slightly asymmetric and relatively narrow and relatively wide 8 short and pointed long and pointed long and bluntly pointed short and pointed short and pointed 9 11-16 8-12 8-14 9-12 8-10 10 absent absent absent absent absent 11 rounded wide and rounded wide and rounded very rare, rounded solitary, rounded or bluntly pointed 12 present mainly absent absent present present 13 strongly asymmetric asymmetric slightly asymmetric slightly asymmetric asymmetric 14 absent absent present present present 15 5-9 at least 10 at least 15 9-12 6-9 16 pointed rounded rounded pointed rounded and bluntly pointed 17 3/4 2/3-3/4 2/3-3/4 3/4 2/3-3/4 18 orange to brownish-red brown to dark brown 19 orange to brownish-red, without ring yellowish brown with brown rings 20 castaneous-brown dark brown 21 3 3 22 approximately square slightly elongated 23 with subparallel margins with subparallel margins 24 almost not widening 2/3 segment length 25 slightly concave concave 26 slightly elongated mainly quadrangular or oval

246 R. J. GODUNKO et al. Table 3 CH BIN BC BPR 1 elongated, widened elongated, rounded or bluntly pointed elongated, rounded 2 elongated and wide rounded or pointed small and wide 3 1.45 1.50 1.65 4 1 + 5-10 1 + 5-7 1 + 8-10 5 with one pointed scale with one rudimentary scale with one rudimentary scale 6 3 3 3 7 slightly asymmetric and relatively narrow relatively symmetrical slightly elongated, relatively symmetrical and narrow 8 long and rounded long and rounded long and pointed 9 8-11 9-12 7-9 10 absent present present 11 wide and rounded bluntly pointed bluntly pointed 12 present present present 13 asymmetric slightly asymmetric slightly asymmetric 14 present absent present 15 9-11 at least 15 at least 8 16 rounded pointed pointed 17 1/2-2/3 1/2-1/3 1/8-1/9 18 light yellow to brownish yellowish-orange to brownish-orange 19 yellowish, without ring light orange, without ring 20 brown brown 21 3 3 22 approximately square not square, slightly elongated 23 with mainly subparallel margins and strong inner projection not elongated, with subparallel margins 24 1/2-2/3 segment length 1/2-2/3 segment length 25 clearly concave clearly concave 26 oval oval REFERENCES BRODSKY K. Zur Kenntnis der mittelasiatischen Ephemeropteren I. (Imagines). Zoologische Jahrbücher, 59: 681-720. ENGBLOM E. 1996. Ephemeroptera, Mayflies. [In:] A. NILSSON (ed.) Aquatic Insects of North Europe A Taxonomic Handbook. Apollo Books, Stenstrup: 13-53. FUJITANI T., HIROWATARI T., TANIDA K. 2003. Nymphs of Nigrobaetis, Alainites, Labiobaetis, Tenuibaetis and Baetis from Japan (Ephemeroptera: Baetidae): Diagnoses and keys for genera and species. [In:] E. GAINO (ed.) Research update on Ephemeroptera & Plecoptera. Università di Perugia, Perugia: 127-133 pp.

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248 R. J. GODUNKO et al. TEMIROVA S. I., PARTOLAKHA N. V., TUROBOV A. L. 1984. Zooplankton i makrozoobentos verkhnego techeniya reki Biyuk-Karasu v svyazi s problemoi okhrany malykh rek. Prirodnye kompleksy Kryma, ikh optimizatsiya i okhrana. Izd-vo SGU. Simferopol, 135-141 pp. (In Russian). ZIMMERMANN W. 1980a. Baëtis braaschi n. sp., ein bisher unbekannter Vertreter der rhodani Gruppe von der Krim (UdSSR) (Ephemeroptera, Baëtidae). Reichenbachia, 18: 199-202. ZIMMERMANN W. 1980b. Beitrag zur Kenntnis der Gattung Baetis LEACH, 1815 (Insecta, Ephemeroptera) im Kaukasus und in Transkaukasien (UdSSR). Entomologische Nachrichten, 25(7-8): 97-112.